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JAN 10 1867

Postilla

UNIVERSITY
PEABODY MUSEUM OF NATURAL HISTORY

YALE UNIVERSITY
NEW HAVEN, CONNECTICUT, U.S.A.

Number 104 December 28, 1966

TWO NEW SPECIES OF CLYMENELLA (POLYCHAETA:
MALDANIDAE) FROM BRAZIL

CHARLOTTE P. MANGUM*

DEPARTMENT OF BIOLOGY, COLLEGE OF WILLIAM AND
Mary, WILLIAMSBURG, VIRGINIA

INTRODUCTION

A collection of maldanid polychaetes made by Dr. Liliana
Forneris in Brazil was recently placed at my disposal. The collec-
tion contains three species from shallow waters of the Atlantic
Ocean in the vicinity of the Marine Laboratory of the Instituto
Oceanografico of the Universidade de Sao Paulo at Ubatuba, Bra-
zil (S. Lat. 23° 30’, W. Long. 45°). Descriptions of two new
species and notes on a poorly known third follow below.

GENUS CLYMENELLA VERRILL 1873

Clymenella (= Euclymene) dalesi sp. nov.
SYNONYM. Questionably Praxilla kefersteini Kinberg, 1866.
DESCRIPTION. Fully formed adult with 25 segments: a single

achaetous prostomial segment, 21 setigerous segments, 2 pre-anal
achaetous segments and a single achaetous anal segment.

*Systematics-Ecology Program, Marine Biological Laboratory, Woods Hole,
Mass. SEP Contribution No. 89.

2 Postilla Yale Peabody Museum No. 104

Cephalic plate slanting dorsally on prostomium at an acute
angle (60-80°). Rim of cephalic plate notably elevated, flaring,
conspicuously notched: a pair of anteroventral notches, a pair
of lateral notches and a single median dorsal notch. Rim other-
wise entire, lacking crenulations and denticles. Central depression
of cephalic plate bisected by longitudinal median keel, which is
bounded on either side by a deep straight furrow (nuchal organ)
extending approximately three-quarters the length of the plate.
Keel protruding anteroventrally between rim notches as small
papilla. Prostomium lacking dark pigment granules. Prostomial
bulb irregularly but not conspicuously papillate (Fig. 1A).

First three setigerous segments with bundle of notopodial capil-
lary setae and, typically, a single neuropodial uncinus. Five of the
40 specimens with two uncini per neuropodium on at least one
side of one segment. Capillary setae long (1.5 mm), slender (7),
primarily bilimbate, though some are spinulose towards sharply
pointed tips. No pennate capillary setae clearly noted. Uncini
large (500-700, long, 50u wide), yellow, with vertical and hori-
zontal striations on shaft. Tips smooth, lacking teeth, notches and
beards; tips typically bent at angle of 120°, but range from 100 to
1GOniCHig. Ie).

Fourth setiger with fleshy anterior rim which may project over
posterior portion of third in the contracted state; one of 40 speci-
mens with freely flanged collar. Setigers 9-21 becoming increas-
ingly elongate; parapodia protruding. Posterior notopodial setae
similar to anterior ones; posterior neuropodia with single row of
up to 20 rostrate (angle approximately 30°) uncini. Posterior
neuropodial setae small (150-180 long, 18 wide), bearded,
bulbed at epidermal level; subdermal shaft vertically striated; tips
with 4-5 small teeth plus main fang (Fig. 1D).

Anal funnel with irregularly alternating long and short cirri,
often becoming triangular in large specimens. Relative length of
anal cirri quite variable, but median ventral cirrus never longer
than rest (Fig. 1B). Anus central in depression of funnel.

Preserved specimens colorless to dark brown with discrete red
bands encircling portions of setigers 4-8.

Mean length of 10 complete specimens, 39.6 mm; range,
12-73 mm; width, 2-3 mm.

Tube unknown.

1966 Two New Species of Clymenella

Figure 1. Clymenellla dalesi sp. nov.

A. Head region, lateral view.
B. Tail region, ventral view.
C. Anterior neuropodial seta.
D. Posterior neuropodial seta.

4 Postilla Yale Peabody Museum No. 104

Locaitigs. Bay of Flamengo (depth 1-12 m., salinity 34.12
— 35.34 0/00); Santos.

Discussion. The generic separation of Euclymene Verrill
(1900) from Clymenella Verrill (1873, 1900) is by no means
clearly defined at present. The distinction is sometimes based on
the presence of a flanged collar on the fourth setiger of Clymenella,
although Verrill (1900) himself emphasized the superficial nature
of this character and chose not to regard it as a generic one.
Moreover, at least two of the four species of Clymenella recog-
nized by Hartman (1959) have no collars. Alternatively, the
distinction is sometimes made on the presence of anterior neuro-
podial aciculae (needles or bent spines) in Euclymene as opposed
to rostrate uncini (recurved hooks) in Clymenella. Since the same
difference is found within the genus Praxillella, it is illogical to
accord generic status to this character alone. Finally, the categori-
cal distinction between aciculae and rostrate uncini certainly exag-
gerates the differences by implying a dichotomy where in fact a
continuum exists. The evidence in support of this contention will
be presented in a discussion cf maldanid phylogeny which is in
preparation.

Clymenella dalesi is the only species in either genus which is
known to have 21 setigerous segments. There are, however, six
species recognized by Hartman (1959) whose segment number is
not known. None of these incompletely known species occurs in
parts of the world where gene exchange is likely to be maintained
with a Brazilian population; moreover, morphological characters
alone suffice to distinguish C. dalesi from these six as follows:

Grube (1840) described 24 segments in the Mediterranean
species Clymene palermitana. On re-examination of the type,
Fauvel (1927, p. 176) indicated that it has “20 a 22 sétigéres
(?)” [sic] and three clearly illustrated achaetous pre-anal seg-
ments. Regardless of the correct number of setigers, the three
achaetous pre-anal segments of this species, listed as Euclymene
palermitana by Hartman (1959), distinguish it from the Brazilian
worm. Fauvel (1927) also mentions the occasional presence of
prostomial pigment granules and a longer median ventral cirrus,
both of which are absent in C. dalesi.

Fauvel’s (1927) account of a fragment from the English
Channel, Clymenella (?) cincta (Saint-Joseph), clearly shows a

1966 Two New Species of Clymenella >

festooned, flanged collar on the fourth setiger which is absent in
C. dalesi. In addition, Saint-Joseph (1894) and Fauvel (1927)
both described only anterior and ventrolateral notches in the
cephalic rim — no posterior notch as in C. dalesi. In both cases,
the figures agree with the text.

Euclymene coronata Verrill (1900), from the Bermuda Is-
lands, has only slight lateral notches in the cephalic rim, and 8-10
lobes on the posterior border. Moreover, I have recently examined
two specimens from Tavernier Key, Florida; each has 19 setigers.

Euclymene tropica (Monro, 1928), which consists of both
anterior and posterior fragments from the Pacific coast of Panama,
also has a crenulated, denticulated posterior cephalic rim, which
is clearly illustrated. In addition, it has only one achaetous pre-
anal segment.

Moore (1923) described the two species Euclymene delineata
and E. reticulata from fragments collected off southern California.
E. reticulata is str:kingly distinct from all others in the genus by
virtue of its completely united prostomial and first setigerous seg-
ments. EF. delineata differs from C. dalesi by virtue of its longer
median ventral anal cirrus (Hartman and Barnard, 1960), and
possibly by its toothed posterior cephalic rim. Although Moore
(1923) emphasized the deep reticulation of its integument, cer-
tainly not true of the specimens of C. dalesi at hand, I am skeptical
of the taxonomic value of this character, which is often correlated
with size.

I have mentioned above a highly uncertain synonym, Praxilla
kefersteinit Kinberg (1866), which is considered indeterminable
(Arwidsson, 1907). The few details of Kinberg’s (1866) descrip-
tion do agree with C. dalesi, but they are not diagnostic.

It gives me great pleasure to name this species in honor of my
colleague, Dr. R. Phillips Dales of Bedford College, University of
London.

Ho.LotyPe. Ubatuba, Bay of Flamengo. YPM No. 2596.
PARATYPES. Same locality. USNM No. 34089; YPM Nos.
297 2598:
Clymenella (= Axiothella) brasiliensis sp. nov.

SYNONYM. Questionably /phianissa armata Kinberg (1866).

6 Postilla Yale Peabody Museum No. 104

DEscCRIPTION. Fully formed adult with 22 segments: a single
achaetous prostomial segment, 18 setigerous segments, 2 pre-anal
achaetous segments and a single achaetous anal segment.

Cephalic plate slanting dorsally on prostomium at an acute
angle (approximately 65°). Rim of cephalic plate notably ele-
vated, flaring; rim conspicuously notched: a pair of anteroventral
notches, a pair of lateral notches and a single median dorsal notch;
rim otherwise entire, lacking crenulations and denticles. Central
depression of cephalic plate bisected by median longitudinal keel,
which is bounded on either side by a deep straight furrow (nuchal
organ) extending one-half to two-thirds the length of the plate.
Keel protruding anteroventrally as small papilla. Dark pigment
granules (often called “ocelli” quite erroneously, for photosensi-
tivity is unknown in maldanids) present immediately ventrolateral
to keel papilla, but inconspicuous. Prostomial bulb faintly and
irregularly papillate (Fig. 2A).

First three setigerous segments with notopodial capillary setae
and several neuropodial uncini. Capillary setae long (1.3 mm),
slender (12), either bilimbate with dark vertical striations or
finely pennate towards tips. Spinulose capillary setae not noted.
Neuropodial setae numbering 4-6 on first setiger, 4-6 on second
and 3-5 on third; typically descending in number from first to third
setiger. Neuropodial setae rather large (450-520 long, 25-30p
wide) with dark brown vertical striations on shaft and a distinct
notch (but no beard) immediately proximal to the main fang.
Tips bent at angle of 85-120°, with 1-3 small teeth plus main
fang (Fig. 2C).

Fourth setiger with fleshy anterior rim which may project over
posterior portion of third in contracted state, but no freely flaring
collar in the 11 specimens. Setigers 9-18 becoming increasingly
elongate; parapodia protruding. Posterior notopodial setae similar
to anterior ones; posterior neuropodia with up to 15-25 rostrate
uncini in single row. Uncini small (200-270, long, 18 wide),
bearded, notched at epidermal level; subdermal shaft faintly
striated vertically. Tips bent acutely (angle 15-25°); 4-6 teeth
plus main fang (Fig. 2D).

Anal funnel with irregularly alternating long and short cirri.
Median ventral cirrus 1144 - 2% times longer than others (Fig. 2B).

1966 Two New Species of Clymenella 7

Figure 2. Clymenella brasiliensis sp. nov.

Head region, lateral view.
Tail region, ventral view.

Anterior neuropodial seta.
Posterior neuropodial seta.

com>

Preserved specimens colorless or dark brown, with red bands
encircling portions of setigers 3-9.
Length of 4 complete specimens, 52-64 mm; width, 2-3 mm.

8 Postilla Yale Peabody Museum No. 104

Loca.ities. Bay of Flamengo (depth 1-12 m., salinity 34.12
— 35.34 0700); Santos.

Discussion. Verrill (1900) explicitly relegated Axiothella to
subgeneric status, giving his reasons at some length. I have re-
viewed the history of the arguments previously (Mangum, 1962).

The characters that distinguish C. brasiliensis from other mem-
bers of the genus are the combination of 18 setigerous and 2
pre-anal achaetous segments, the presence of a fleshy rim on the
fourth (and not the fifth) setiger, the presence of a longer median
ventral anal cirrus and the number of cephalic rim notches. It is
also distinguished from Clymenella minor Arwidsson (which does
have the above combination) primarily by the character of the
anterior neuropodial setae, which are virtually straight and unbent,
fewer and larger in C. minor (Arwidsson, 1911). Of perhaps some
importance, C. minor differs in its unique pattern of pigmentation
and the relative length of anal cirri.

Again, I have mentioned above an equally uncertain synonymy
with Iphianissa armata Kinberg (1866), because the few details
given do agree with C. brasiliensis. Arwidsson’s (1907) referral
of [phianissa Kinberg to Praxillella Verrill, on the basis of similar
neuropodial setae and the location of glands, is not valid since
these characters alone do not distinguish Praxillella from at least
three other genera.

Ho.otyPe. Ubatuba, Bay of Flamengo. YPM No. 2593.

PARATYPES. Same locality. USNM No. 34090; YPM Nos.
2594, 2595.

GENUS ASYCHIS KINBERG 1866

Asychis amoena (KINBERG) 1866

DESCRIPTION. Body of 21-22 segments: a single achaetous
prostomial segment, 19 setigerous segments, a single pre-anal
achaetous segment (questionably) and a single achaetous anal
segment.

Cephalic plate slanting dorsally on prostomial segment at acute
angle of approximately 50°. Rim of cephalic plate moderately
elevated, with deep lateral notches on each side, faint ventro-
lateral notches on each side and several irregular faint notches on
median ventral portion. Cephalic plate with pair of inverted

1966 Two New Species of Clymenella 9

U-shaped nuchal organs anteroventrally and incomplete horizontal
furrow posterodorsally. No raised median cephalic keel.

First setigerous segment with laterally notched, flanged collar.
Collar with single, deep scallop beginning mid-dorsally and ending
at right lateral notch.

First setiger with dorsal capillary setae in single row; no ventral
setae. Second setiger with row of 7 neuropodial rostrate uncini
and elliptical fascicle of dorsal capillary setae. Notopodial setae
continuing as ellipse posteriorly; neuropodial setae increasing in
number posteriorly to about 20-25.

Anus dorsal to suboval caudal plate. Rim of caudal plate with
deep ventrolateral notches, otherwise entire. No caudal cirri.
Caudal plate with slight ventral depression, otherwise somewhat
convex.

Preserved specimen essentially colorless, with a very few tiny
dark pigment granules scattered over anterior part of dorsum.
Anterior segments opaque; mid-region and posterior segments
transparent, showing eggs in coelom.

Length of one complete specimen, 43 mm; width, 3 mm.
Tube unknown.

LocALITY. Vitoria Island.

Discussion. Although the original description of this species
is rather sketchy (Kinberg, 1866), Hartman (1949) rede-
scribed the type material in full detail. The type material con-
sists of a single complete specimen from the shallow (43-66 m)
waters of the Atlantic Ocean about 650 km northeast of Vitoria
Island. The present collection also includes a single specimen
which agrees in most respects with the descriptions given by
Kinberg (1866) and Hartman (1949). It is difficult to believe that
the most conspicuous discrepancy, the asymmetrical scallop on the
collar of the first setiger, is truly typical, although there is no evi-
dence of damage to the specimen. It should probably be regarded
as an abnormal growth in this individual.

Since the species is known only from the type material, the
description above may contribute to our knowledge of the range
of variation found within it.

I have donated my single specimen to the Peabody Museum
at Yale University (YPM No. 2592).

10 Postilla Yale Peabody Museum No. 104

ACKNOWLEDGMENTS

I am indebted to Mrs. Ruth von Arx, who prepared the illustra-
tions, and to Drs. Meredith L. Jones and J. L. Simon for criticisms
of the manuscript. Drs. Alyton Joly and Edmondo Nonato and
Messrs. Manoel Marcelino and Clarimundo de Jesus assisted in
the collection and preparation of specimens. The work was sup-
ported in part by a grant from the Whitehall Foundation to the
Systematics-Ecology Program, Marine Biological Laboratory,
Woods Hole, Mass.

LITERATURE CITED

Arwidsson, I. 1907. Studien tiber die skandinavischen und arktischen Mal-
daniden nebst Zusammenstellung der iibrigen bisher bekannten Arten
dieser Familie. Zool. Jahrb., Suppl. 9: 1-308.

1911. Maldaniden. Wiss. Ergebn. Schwed. Sudpolar-exped. 6:
1-44.

Fauvel, P. 1927. Faune de France. 16. Polychetes sédentaires. Lechevalier.
Paris. 494 p.

Grube, A. E. 1840. Actinien, Echinodermen und Wiirmer des Adriatischen
und Mittelmeers. J. H. Bon. Konigsberg. 92 p.

Hartman, O. 1949. The marine annelids erected by Kinberg with notes on
some types in the Swedish State Museum. Ark. f. Zool. 42: 1-137.

1959. Catalogue of the polychaetous annelids of the world.
Allan Hancock Found., Occ. Pap. 23. 628 p.

Hartman, O., and J. L. Barnard. 1960. The benthic fauna of the deep basins
off southern California. Allan Hancock Pacific Exped. 22 (2). 297 p.

Kinberg, J. G. H. 1866. Annulata nova. Ofvers. K. VetenskAkad. Forh. 23:
337-357.

Mangum, C. P. 1962. Studies on speciation in maldanid polychaetes of
the North American Atlantic coast. I. A taxonomic revision of three
species of the subfamily Euclymeninae. Postilla (Peabody Museum,
Yale Univ.) 65. 12 p.

Monro, C.C.A. 1928. On the Polychaeta collected by Dr. Th. Mortensen
off the coast of Panama. Vidensk. Meddr. Dansk Naturh. Foren. 85:
75-103.

Moore, J. P. 1923. The polychaetous annelids dredged by the USS “Alba-
tross” off the coast of southern California in 1904. IV. Spionidae to
Sabellariidae. Acad. Nat. Sci. Philadelphia, Proc. 75: 179-259.

Saint-Joseph, Baron de. 1894. Les annélides polychétes des cotes de Dinard.
Sci Nat... Ann® (7erser)) 17e 1-395:

Verrill, A. E. 1873. Report on the Invertebrata of Vineyard Sound and
adjacent waters, with an account of the physical characters of the
region. U.S. Comm. Fish and Fisheries for 1871-72, Rep.: 295-852.

—— 1900. Additions to the Turbellaria, Nemertina, and Annelida
of the Bermudas, with revisions of some New England genera and
species. Conn. Acad. Arts & Sci., Trans. 10: 595-698.

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